Received: 10 March 2023 Accepted: 2 January 2024

DOI: 10.1002/agj2.21539

REVIEW
Crop Econom ics, Production, and Managem ent

Sustainable sweetpotato production in the United States: Current

status, challenges, and opportunities
Justin George1 Gadi V. P. Reddy1 Phillip A. Wadl2 William Rutter2
Julianna Culbreath2 Pierre W. Lau3 Tahir Rashid4 Matthew C. Allan5
Suzanne D. Johaningsmeier5 Amanda M. Nelson6 Ming Li Wang7
Augustine Gubba8 Kai-Shu Ling2 Yan Meng9 Daniel J. Collins9
Sathish K. Ponniah10 Prasanna H. Gowda11
1 Southern Insect Management Research Unit, USDA-ARS, Stoneville, Mississippi, USA
2 U. S. Vegetable Laboratory, USDA-ARS, Charleston, South Carolina, USA
3 Pollinator Health in Southern Crop Ecosystem Research Unit, USDA-ARS, Stoneville, Mississippi, USA
4 Extension/Research Demonstration Farm & Technology Transfer Center, Alcorn State University, Mound Bayou, Mississippi, USA
5 Food Science and Market Quality and Handling Research Unit, USDA-ARS, Raleigh, North Carolina, USA
6 National Center for Alluvial Aquifer Research, USDA-ARS, Stoneville, Mississippi, USA
7 Plant Genetic Resources Conservation Unit, USDA-ARS, Griffin, Georgia, USA
8 Discipline of Plant Pathology, School of Agricultural, Earth and Environmental Sciences, University of Kwazulu-Natal, Pietermaritzburg, South Africa
9 Department of Agriculture, Alcorn State University, Alcorn, Mississippi, USA
10 Department of Agriculture, University of Arkansas at Pine Bluff, Pine Bluff, Arkansas, USA
11 AJWDSRC, USDA-ARS Southeast Area, Stoneville, Mississippi, USA

Correspondence
Justin George, Southern Insect Management
Research Unit, USDA-ARS, 141
Experiment Station Rd., Stoneville, MS
38776, USA.
Email: justin.george@usda.gov
Assigned to Associate Editor Nothabo
Dube.
Funding information
United States Department of Agriculture,
Agricultural Research Service, Grant/Award
Number: 6066-22000-090-00D
Abstract
Sweetpotato (Ipomoea batatas L.) is an important staple crop cultivated in over 100
countries, and the storage roots and vines provide food for humans and livestock.
Sweetpotato consumption and demand for its value-added products have increased
significantly in the last two decades and have led to new cultivar development, expan-
sion in acreage, and increased demand in the United States and its export markets.
Despite the known nutritional components and other health benefits, further research
is needed to characterize the genetic diversity and chemical composition related
to their storage root qualities, essential in developing consumer-preferred cultivars
that offer host plant resistance against pests and pathogens. There is a critical need
for research on non-pesticidal control approaches that can provide safe, effective,
economical, sustainable, and environmentally sound pest and disease management
techniques, especially for socially disadvantaged small farmers in the United States.
Moreover, climate change can significantly impact future production practices and

Abbreviations: CIP, international potato center; IPM, integrated pest management; PPN, plant parasitic nematode; RKN, root-knot nematode; RN, reniform
nematode; WU, water use; WUE, water use efficiency.

This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any medium, provided
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© 2024 The Authors. Agronomy Journal published by Wiley Periodicals LLC on behalf of American Society of Agronomy. This article has been contributed to by U.S. Government
employees and their work is in the public domain in the USA.

Agronomy Journal. 2024;1–31. wileyonlinelibrary.com/journal/agj2 1

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2 GEORGE ET AL.

yield and may directly or indirectly affect crop pests, weeds, and diseases. In this
review, we discuss the current status, challenges, and future approaches associated
with sweetpotato production practices; health-promoting properties of sweetpotato
cultivars; value-added products; genetic diversity and germplasm; pest and disease
management; weed and water management; pollination ecology; and other agro-
nomic and cultural practices that may impact sustainable sweetpotato production by
small-scale, organic, and large-scale growers.

1 INTRODUCTION
Sweetpotato, Ipomoea batatas L., is a favorite staple crop
in many cultures and is established worldwide (Mukhopad-
hyay et al., 2011; Woolfe, 1992). It is a dicotyledonous plant
belonging to the family Convolvulaceae and a root crop
important for food security (Barkessa, 2018). Worldwide,
sweetpotato is the seventh most important food crop after
rice, wheat, potato, maize, barley, and cassava (Muhammad
et al., 2012; Neela & Fanta, 2019). From 2018 to 2021, the
estimated global production ranged from 88.7 to 92.3 Mt,
where Asian countries produced 61.5%–66.6% and African
countries 28.6%–33.7% of the global production (FAOSTAT,
2023). Sweetpotato is a valuable crop because it is regarded
as one of the most nutritious vegetables and produces more
food per hectare than any other crop. It is also a versatile
crop because the storage roots can be consumed or processed
into value-added foods (e.g., fries, chips, starch, alcohol) and
industrial products (e.g., fuel and chemicals) plus the vines
can be consumed or used as livestock feed (Clark et al., 2012;
Loebenstein & Thottappilly, 2009; Zhang et al., 2013).
North Carolina is the leading sweetpotato producer by
tonnage in the United States, followed by California and Mis-
sissippi (USDA-NASS, 2023). Despite the increase in acreage
and cultivars, the sweetpotato industry faces many challenges
associated with yield losses due to weather extremes, pest and
disease incidence, storage and processing issues, and under-
use of value-added products. Sweetpotato production and the
number of farmers (small, large, conventional, and organic)
interested in sweetpotato cultivation are growing. However,
the economic viability of sweetpotato production can be
undermined due to susceptibility of commercial cultivars
to biotic (i.e., diseases, insect pests, and weeds) and abi-
otic (i.e., environmental variability, low fertility) factors that
can dramatically influence yield, quality, and marketability.
Management practices such as pest and disease management
for sweetpotato can be very expensive, add substantially to
market prices, and have adverse environmental and health
effects. Socially disadvantaged small farmers are more vul-
nerable to losses due to a lack of integrated pest management
(IPM) knowledge, limited resources, and challenges in man-
aging plant pests, as most IPM projects focus on large farms
(Collins, 2022). Hence, there is a critical need for research
on nonchemical control approaches that can provide effective,
sustainable, and environmentally sound pest and disease man-
agement techniques for socially disadvantaged small farmers
in the United States. Nonetheless, sweetpotato is an ideal crop
for limited resource farmers as it produces more biomass and
nutrients per hectare than any other food crop globally without
fertilizers and irrigation (Loebenstein & Thottappilly, 2009)
and host resistance to the common plant pests is readily avail-
able (Jones et al., 1986). The current status, challenges, and
solutions for the US sweetpotato industry need to be identified
and discussed to support sweetpotato cultivation.
This is an extensive review on the US sweetpotato indus-
try from subject matter experts to address these needs. The
topics that will be addressed are (1) health-promoting prop-
erties of sweetpotato cultivars, value-added products, and
consumer preferences; (2) pollination ecology and the forag-
ing landscape in sweetpotato ecosystems; (3) pest and disease
management; (4) weed management; (5) water management;
(6) impact of climate change on global sweetpotato produc-
tion; and (7) current limitations and challenges for small-scale
and organic farmers. Also, we discuss how these agronomic
and cultural practices impact sustainable production by small-
scale, organic, and large-scale growers (Figure 1). These
topics impact all US sweetpotato industry stakeholders and
demonstrate the need for increased research efforts to support
this industry.
2 HEALTH-PROMOTING PROPERTIES
OF SWEETPOTATO CULTIVARS AND
CONSUMER ACCEPTANCE
2.1 Health benefits of sweetpotatoes
The sweetpotato is a nutritious vegetable with substantial
quantities of essential vitamins and minerals and health-
promoting phytonutrients. For example, a baked medium-
sized sweetpotato (114 g) is high in copper, manganese, and
vitamins A, C, B1, B2, B3, B5, and plus B6 and is a good

GEORGE ET AL.
F I G U R E 1 Interactions of different factors that may contribute to
the sustainable production and consumption of sweetpotatoes. All these
factors and changes in weather patterns can influence the sweetpotato
production, quality, and yield by small-scale, organic, and large-scale
producers.
source of potassium (USDA-ARS, 2019). Sweetpotatoes are
also a good source of fiber, which has been shown to improve
the gut microbiome and promote beneficial immunological
responses (Liu et al., 2020; Tang et al., 2018).
Sweetpotatoes flesh can be colored orange, yellow, white,
purple, and shades in between. Orange-fleshed sweetpota-
toes garner their color from β-carotene—a carotenoid with
antioxidant and pro-vitamin A activities. About 90% of the
carotenoids in Covington, a popular orange-fleshed culti-
var in the United States, is β-carotene (Grace et al., 2014).
β-Carotene is a unique carotenoid because it has 100% pro-
vitamin A activity, and one molecule can be converted into
two vitamin A molecules in the body. This makes orange-
fleshed sweetpotatoes exceptionally nutritious, particularly in
regions suffering from vitamin A deficiencies (Boy & Miloff,
2009). Yellow-fleshed sweetpotatoes also contain carotenoids
but in lower concentrations with ~30%–60% being β-carotene
(Grace et al., 2014). Carotenoids in the diet reduce metabolic
oxidative stress, are beneficial for the immune system, and
reduce cancer and cardiovascular risks (Rodriguez-Amaya,
2015).
The color of purple sweetpotatoes is due to anthocyanins,
a class of pH-sensitive, water-soluble compounds commonly
found in red-, blue-, and purple-colored fruits and vegeta-
bles (Wu et al., 2006). Anthocyanins are polyphenols with
powerful antioxidant activities and are generally associated
with health-promoting benefits. Purple sweetpotatoes and
extracts thereof have been associated with antioxidant activ-
ity, anticarcinogenic effects, anti-inflammatory properties,
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3
Core Ideas
∙ Sweetpotato (Ipomoea batatas L.) is an important
staple crop cultivated in over 100 countries.
∙ US sweetpotato industry faces many production
challenges, including pest and diseases, as well as
climate change extremes.
∙ A comprehensive review by subject matter experts
on the challenges of US sweetpotato industry is not
available.
∙ This review evaluates the current situation, chal-
lenges, and future approaches for improving sweet-
potato production.
∙ Also, current and future impacts of climate change
on global sweetpotato production and demand are
discussed.
blood glucose modulation, immunomodulatory activity, plus
liver and kidney protection (Albuquerque et al., 2019; Jiang
et al., 2022). Purple sweetpotato anthocyanin content can
vary, but Stokes Purple, one of the leading purple cultivars
in the United States, has similar antioxidant levels on a fresh
weight basis as antioxidant-rich fruits, such as grapes, straw-
berries, and raspberries (Truong et al., 2010; Wu et al., 2006).
In addition to differences in total anthocyanin concentra-
tion, the types of anthocyanins also affect purple sweetpotato
bioactivities (Esatbeyoglu et al., 2017; Hu et al., 2016). There
is a wealth of data demonstrating sweetpotatoes are a healthy
food (Albuquerque et al., 2019). Therefore, developing new
varieties and technologies that increase sweetpotato consump-
tion in the United States would benefit both the sweetpotato
industry and the health of the American people.
2.2 Common US sweetpotato cultivars
The most popular type of sweetpotatoes in the United States
are the sweet, moist, light-skinned, and orange-fleshed culti-
vars Covington and Beauregard (Table 1). In some parts of the
country, these varieties are marketed as “yams,” a term intro-
duced by southern US sweetpotato producers to distinguish
orange-fleshed cultivars from the white/cream-fleshed vari-
eties of the mid-20th century. To avoid confusion with the true
yam (Dioscoreaceae family), the U.S. Department of Agri-
culture (USDA) now requires that the word “sweetpotato” is
present on the label (Smith et al., 2009; Truong et al., 2018).
Covington and Beauregard are the current dominant cultivars
of this market category, but Orleans is a newer orange-
fleshed cultivar that has greater yields than Beauregard (La
Bonte et al., 2012) and is starting to capture a portion of the
 4

TA B L E 1 Characteristics and compositions of commonly grown US sweetpotato cultivars.

Dry Starch Sucrose Glucose + fructose β-Carotene Monomeric anthocyanins

Cultivar Flesh colora Skin colora n matter % (g·100 g−1 FW) (g·100 g−1 FW) (g·100 g−1 FW) (μg·g−1 FW) (μg·g−1 FW)
Bayou Belle Orange Red/purple 76 20.6 ± 1.3 6.5 ± 1.0 2.4 ± 0.4 2.2 ± 0.4 69.4 ± 18.2 ND
Beauregard Orange Light rose 205 21.9 ± 1.7 9.5 ± 1.4 1.5 ± 0.4 1.6 ± 0.4 72.9 ± 19.5 ND
Covington Orange Light rose 263 22.2 ± 1.5 7.8 ± 1.4 2.9 ± 0.4 1.0 ± 0.3 71.6 ± 19.1 ND
Jewel Orange Copper 8 22.7 ± 2.2 9.2 ± 2.2 1.8 ± 0.3 1.4 ± 0.3 73.1 ± 15.8 ND
Orleans Orange Light rose 12 21.1 ± 1.3 9.2 ± 1.1 1.4 ± 0.3 1.7 ± 0.2 70.7 ± 20.2 ND
Murasaki-29 White Dark purple 28 28.1 ± 1.2 16.0 ± 1.9 2.1 ± 0.2 0.4 ± 0.2 ND ND
Bonita White/cream Light tan 23 26.3 ± 1.3 14.3 ± 2.0 1.6 ± 0.3 0.8 ± 0.2 ND ND
Purple Dark purple Dark purple 41 28.5 ± 1.0 15.3 ± 1.4 1.3 ± 0.4 1.1 ± 0.3 –b 773.0 ± 95.2
Majesty
Purple Purple Dark purple 57 28.3 ± 1.0 16.0 ± 1.6 1.8 ± 0.2 0.6 ± 0.2 –b 627.0 ± 114.9
Splendor
Stokes Purple Dark purple 39 28.6 ± 1.3 17.5 ± 2.0 1.5 ± 0.3 0.4 ± 0.2 –b 498.5 ± 160.5
Purple
Note: Compositions are predictions from near-infrared spectroscopy (NIRS) scans of roots that were stored 6–10 weeks after harvest from 2017 to 2021. Data provided by the North Carolina State University Sweetpotato and Potato
Breeding and Genetics Programs.
Abbreviations: FW, fresh weight basis; ND, not detected.
a
Sweetpotato characteristic descriptions from LSU and NCSU sweetpotato breeding programs (Arnold, 2016; Sweetpotato and Potato Breeding and Genetics Programs, 2022).
b
β-Carotene cannot be predicted in purple-fleshed sweetpotatoes by NIRS using the current models.
GEORGE ET AL.

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GEORGE ET AL.
market. Vermillion and Diane are also orange-fleshed cul-
tivars but have thicker red-purple skin and are sometimes
labeled as “red-yams” or “Garnet” at the point of sale. Bayou
Belle is another high-yielding cultivar in this market category
but is mainly grown for processing (Table 1). The “red-yam”
is prevalent in California and often receives higher prices than
the “yam” sweetpotatoes (Stoddard et al., 2013).
Cream/yellow-fleshed sweetpotatoes with light skin tend to
have higher dry matter and a less moist texture than orange-
fleshed cultivars. They are marketed as “Jersey Sweets”
particularly in the Northeastern United States and “sweets”
or “sweetpotatoes” in the western states and provinces of the
United States and Canada (Smith et al., 2009; Stoddard et al.,
2013). Most of these sweetpotatoes are grown in California
and typically sold at West Coast retailers, high-end grocers,
and ethnic markets (Don La Bonte, personal communication,
2023). Bonita is a popular cultivar in this category, replacing
O’Henry, a cream-fleshed mutant of Beauregard. The “Orien-
tal”/“Japanese” sweetpotato market types have white/cream
flesh with dark purple skin, and Murasaki-29 is the domi-
nant cultivar of this type in the United States (Stoddard et al.,
2013).
Purple sweetpotatoes are currently a niche market in the
United States, and the most common cultivars are Stokes
Purple, with deep purple-flesh and skin, and Okinawan, a
light purple-flesh and light brown skin cultivar. However,
yields of these varieties are less than other sweetpotatoes.
The North Carolina State University sweetpotato breeding
program released two purple-fleshed sweetpotato cultivars in
2021—Purple Majesty and Purple Splendor (Sweetpotato and
Potato Breeding and Genetics Programs, 2022). These cul-
tivars have superior yields, shapes, and growing condition
adaptability with eating quality similar to Stokes Purple (Yen-
cho & Pecota, 2022a, 2022b). Purple sweetpotatoes tend to
have a dry texture and unique flavor characteristics such as
vanilla aroma and more bitter and umami tastes, which were
objectionable in a North Carolina consumer acceptance study
(Leksrisompong et al., 2012). Despite purple sweetpotatoes
having a niche market, there is still a need to develop pur-
ple sweetpotato cultivars that are more favorable to the US
population. This would require an understanding of which
compounds are health promoting as well as those inducing
undesirable flavors; thus, more food science and breeding
research is needed for the development of healthy and tasty
purple sweetpotatoes.
2.3 Sweetpotato eating quality
Consumer preferences for sweetpotato vary within the United
States and around the world. The most popular type in the
United States is the sweet, moist, orange-fleshed sweetpotato
(Barkley et al., 2017; Leksrisompong et al., 2012; Smith et al.,
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5
2009), particularly in the Southeastern United States where
most of the sweetpotatoes are produced. In the Western United
States, the drier, yellow to white-fleshed varieties, along with
moist-orange varieties, are preferred, whereas in Hawaii, dry,
purple-fleshed varieties are preferred (Miyasaka et al., 2019;
Smith et al., 2009; Stoddard et al., 2013). Differences in con-
sumer preferences are also evident in other countries. For
example, moist, orange-fleshed sweetpotatoes were disliked
in South African and Ugandan consumer studies, while the
white, sweet, dry/firm varieties were preferred (Laurie et al.,
2013; Mwanga et al., 2020). Notwithstanding these general
preferences, distinct market segments exist for sweetpota-
toes, each with preferences for different sensory traits (Dery
et al., 2021; Leksrisompong et al., 2012). This provides an
opportunity to expand the overall market for sweetpotato by
developing new cultivars and products that appeal to varied
consumer segments. To better understand regional differences
in consumer preference, a universal sweetpotato lexicon was
developed to objectively evaluate sensory attributes (Nakitto
et al., 2022). This lexicon was further developed to investi-
gate the impact of intrinsic sweetpotato properties on eating
quality of a wide range of US genotypes (Johanningsmeier
et al., unpublished). This work will help breeders, growers,
and processors make cultivar selections that match their mar-
kets’ preferences and develop new markets through varietal
diversification.
2.4 Processed sweetpotatoes: Value-added
products
In the United States, sweetpotatoes that are sold to the con-
sumer are typically US No. 1 grade, which are storage roots
that have a uniform shape, are free of damage, and range in
size between 5.1 and 8.9 cm in diameter and between 7.6
and 22.9 cm in length. Roots that are free from significant
injuries (e.g., decay, cuts, and freezing) but exceed the defect
and irregularity shape limit of the US No. 1 grade can be
graded as US No. 2 or processor grades. Roots larger than
the No. 1 criteria are called “jumbos.” Smaller roots that are
2.5–5.1 cm in diameter and 5.1–17.8 cm in length are called
“canners.” The non-US No. 1 grade sweetpotatoes have lower
value, and the high cost of harvesting can cause a substantial
portion of potentially marketable sweetpotatoes to be unhar-
vested. For example, it was estimated that 125 million kg
of marketable sweetpotatoes were unharvested in the fields
in North Carolina during the 2016 season (Johnson et al.,
2018). Processed sweetpotato products are an avenue for value
addition and utilization of off-grade sweetpotatoes. Common
value-added sweetpotato products in the United States include
French fries, chips, purées, dehydrated products, and canned
forms. Other sweetpotato utilization options include livestock
feed, flour, sugar, and starch production, which are more

6 GEORGE ET AL.
common in other sweetpotato-growing countries (Loeben-
stein & Thottappilly, 2009).
One of the most popular value-added products is sweet-
potato fries. The popularity of this product has grown
substantially over the past two decades, as indicated by
the ~10- to 20-fold increase in the number of times the
term “sweetpotato fries” was searched on Google in 2022
compared to 2004 (Google Trends, 2022). Sweetpotato fry
production is also a great utilization of jumbos and roots with
exterior visual defects. However, fries made from the cur-
rent table-stock varieties lack a crispy exterior and typically
require a batter; as with other foods cooked at high tempera-
tures, they may contain acrylamide, a potentially carcinogenic
neurotoxin (Zyzak et al., 2003). Food science and breeding
research has been conducted to develop processes and geno-
types that are better suited for sweetpotato fries to improve
textures and minimize acrylamide formation. It was identified
that sweetpotato composition (e.g., starch, moisture, sugar
contents), amylase activities, as well as starch viscoelastic,
structural, and thermal properties all affect fry textures (Allan
& Johanningsmeier, 2022; Allan et al., 2021; Sato et al.,
2018). In addition, acrylamide formation can be minimized
in sweetpotato fries by processing aids that interfere with the
reaction or by removal of acrylamide precursors—reducing
sugars and free asparagine (Truong et al., 2014). A rapid
and sensitive liquid chromatography with mass spectrome-
try (LC–MS/MS) method to quantify the limiting substrates
for acrylamide formation has been developed for sweetpota-
toes (Qiu et al., 2020). These findings plus ongoing research
will help develop sweetpotato varieties better suited for fry
production.
Sweetpotato chips are another popular value-added prod-
uct in the United States and are also frequently produced
using the lower starch, higher sugar, orange-fleshed cultivars.
This can lead to challenges with acrylamide formation, exces-
sive browning, weaker textures, and increased oil uptake.
Sweetpotato chip-breaking force and oil contents have been
correlated with dry matter content (Gao et al., 2014; Hagen-
imana et al., 1998), yet dry matter does not fully explain
chip texture or oil content. Sweetpotato chips treated with
enzymes that modified the cell wall polymers also experi-
enced impacts on chip-breaking forces and oil uptake (Allan
& Johanningsmeier, 2022). Therefore, both sweetpotato com-
position and cell wall polymer attributes should be considered
in the selection of genotypes best suited for sweetpotato chips.
Sweetpotato purée is a value-added product that can use
any shape or size of sweetpotato. It is also a versatile prod-
uct that can be used as a biofortifying, healthy ingredient
in many foods (e.g., baby food, beverages, soups, and baked
goods). Sweetpotato purée is a low-acid food (pH 5.8–6.3)
that requires intensive thermal processing for shelf stabil-
ity when using traditional conductive heat transfer—causing
notable nutrient and quality deterioration (Truong et al.,
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2018). However, a continuous flow microwave aseptic pro-
cess developed by USDA-ARS and North Carolina State
University scientists can reach the necessary internal tem-
perature quickly and uniformly. This results in a shelf-stable
product with similar color and viscosity as an unsteril-
ized purée (Coronel et al., 2006; Simunovic et al., 2014).
This advanced sterilization technology enabled the startup of
Yamco, a sweetpotato purée microwave processing facility
located in North Carolina, and an implementation in Kenya
to produce orange-fleshed sweetpotato purées for biofortifi-
cation of baked goods and other foods (North Carolina State
University, 2020).
Sweetpotato dehydration is a value-added process that
results in a shelf-stable product that can be used as an ingre-
dient, ground into flour, or used for the pet food industry
(Boyette & Macialek, 2012; Van Hal, 2000). Depending on
the sweetpotato form (e.g., slices, dices, purée), sweetpota-
toes can be dried using tunnel, oven, drum, or spray dryers
(Truong et al., 2018). Solar drying is another standard method
in rural sweetpotato-growing regions worldwide. However,
a challenge with dehydrated orange-fleshed sweetpotatoes
is β-carotene degradation, with most of it degrading within
the first month of storage. This results in color and nutri-
ent loss and generates off-flavors (Bechoff et al., 2011).
Degradation rates can be slowed by blanching (De Moura
et al., 2015), oxygen removal (Emenhiser et al., 1999), and
antioxidant treatments (Bechoff et al., 2011). However, these
technologies have not been widely implemented in the United
States. Therefore, more research is needed to develop sustain-
able technologies for improving β-carotene retention in dried
sweetpotatoes.
Canned sweetpotatoes are a classic value-added product,
providing a use for smaller roots called “canners” and roots
with visual defects. Peeled roots can be cut into any dimen-
sion or left whole and are typically canned with sugar syrup
(Truong et al., 2018). The high-temperature thermal pro-
cess and long cooling time can cause extensive softening,
likely due to pectin breakdown (Truong et al., 1998). Canned
sweetpotato firmness can be increased by a low-temperature
blanch (62˚C) prior to canning to promote pectin methyl
esterase activity and de-esterification of pectin (Walter et al.,
2003) or by adding sugar and calcium to the canning solution
(Bouwkamp, 1985).
Sweetpotatoes and its products have grown to be a
notable market in the United States, worth around $600–
700 million annually (USDA-NASS, 2023). Continued food
science research in collaboration with breeding and agron-
omy efforts will be needed to sustain growth. This would
include additional comprehensive consumer studies in US
markets coupled with sensory profiling to develop targets
for consumer-preferred sweetpotato cultivars; development of
new technologies to bring additional value to off-grade sweet-
potatoes; and identification of the chemical makeup that is

GEORGE ET AL.
FIGURE 2 The components for the utilization of sweetpotato germplasm collections.
responsible for sweetpotato characteristics that are favorable
for specific processed products.
2.5 Sweetpotato germplasm collection
Sweetpotato germplasm collections contain natural variation,
which provides the genetic base for cultivar development and
improvement. The largest gene bank of sweetpotato is main-
tained at the International Potato Center (CIP) in Lima, Peru,
which holds about 7000 accessions (6000 cultivated acces-
sions and 1000 of its wild relatives) (Anglin et al., 2021).
The USDA-ARS sweetpotato germplasm collection and its
crop wild relatives are maintained at Griffin, GA, by the Plant
Genetic Resources Conservation Unit. This collection con-
tains 606 in vitro clonal accessions and 461 accessions of wild
Ipomoea species maintained as seeds.
There are five major components for any plant germplasm
collection that are interconnected, but the final purpose is
utilization (Figure 2). The highly efficient and maximized uti-
lization of the germplasm collection depends on how to carry
out the tasks of all these five components.
Curation: The number of accessions in each gene bank
is essential, but its number may not reflect its uniqueness.
The passport data (a basic description of an accession) show
redundant accessions from free exchanges that should be iden-
tified. After identifying redundant accessions, sweetpotato
scientists can focus on investigating the unique accessions
for their specific purpose. For any gene bank, there are still
genetic gaps (i.e., existing genetic materials not being col-
lected yet). More accessions should be collected based on
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7
geographic information and country origins to expand genetic
diversity if possible.
Preservation: Compared with other species, the sweet-
potato germplasm accessions are preserved (or maintained)
as clonal materials instead of seeds. Currently, two big chal-
lenges exist for using traditional tissue culture to maintain
sweetpotato germplasm. Because tissue cultures must be sub-
cultured every 3–4 months to preserve cultures, the high
frequency of propagation events can introduce both pathogen
(bacteria, fungi, and virus) infection and somatic mutations.
To prevent losing clonal accessions, cryopreservation meth-
ods provide a highly efficient long-term maintenance of
sweetpotato germplasm (Park & Kim, 2015; Wilms et al.,
2020).
Characterization: Characterization can cover many
aspects, from morphological observation (such as a leaf,
stem, root skin, and flesh color) to agronomical traits (such
as growth rate and maturity) to responses for biotic (such
as resistance to pathogen and pest) and abiotic (tolerance to
drought and severe weather condition) stresses to chemical
composition analysis (such as leaf and root macronutri-
ent and micronutrients). Characterizing specific traits for
research will focus on or utilize a subset of the germplasm
collection. How this subset is selected will directly affect the
characterization results. Using the core (representing 10% of
the entire collection) or mini core (representing 1% of the
entire collection) will be an efficient approach to characterize
the germplasm collection for specific traits (Brown, 1989a,
1989b; Upadhyaya & Ortiz, 2001).
Evaluation: Sweetpotato breeding programs depend on
the germplasm collection to improve nutritional quality,
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8 GEORGE ET AL.

T A B L E 2 Distribution of sweetpotato accessions within the US accessions within the restricted growing areas to prevent them
states during 2020–2021. from becoming out of control.
State Accessions There are different limitations associated with the current
Alabama 12
germplasm collection. The genetic diversity and the chemical
composition related to storage root quality are not well char-
California 45
acterized, which needs further research. Also, the germplasm
Colorado 15
accessions undergo deterioration from somatic mutation over
Georgia 15
multiple generations and require long-term preservation tech-
Indiana 1
niques (such as cryopreservation) to maintain the germplasm
Louisiana 1 collections. New cultivars and breeding lines need to be
Maryland 1 introduced to the collection to expand genetic diversity and
Minnesota 2 selection choices for breeding programs.
North Carolina 1
Ohio 59
Oklahoma 5 3 POLLINATION ECOLOGY AND THE
South Carolina 258 FORAGING LANDSCAPE IN
Washington 17 SWEETPOTATO ECOSYSTEMS
West Virginia 1
A significant portion of large-scale row crop agricultural sys-
Wyoming 2
tems is considered pollinator independent, as they do not
Total accessions 435
require animal-mediated pollination services for the crop to
set fruit. However, pollinators may still visit crops to collect
resources. Bees are known to collect pollen from corn, nectar
from cotton, and pollen and nectar from soybeans (Esquivel
disease/pest resistance, and other agronomic traits. To bet- et al., 2021; Lin et al., 2022). These crops bloom after the
ter utilize the sweetpotato germplasm collections, researchers spring resource flow, providing a valuable option for bees to
have characterized the genetic diversity of global collections obtain valuable nutrients, such as proteins and lipids in pollen
with various DNA marker platforms (AFLP, SLAF, GBSpoly, and carbohydrates in nectar, while other resources are dwin-
chSSR, and SSR) for 97 accessions from Tanzania (Elameen dling as temperatures increase. The presence of pollinators
et al., 2008), 197 accessions from China (Su et al., 2017), and their interactions with crops in these systems have been
303 and 604 accessions from the United States (Slonecki shown to provide benefits to crop yield, providing an oppor-
et al., 2023; Wadl et al., 2018), 558 accessions from Korea tunity for row-crop farmers and beekeepers to coexist with
(Lee at al., 2019), and 5979 accessions from CIP (Anglin the shared interest of expanding their operations (Esquivel
et al., 2021). Based on the number of accessions investi- et al., 2021; Kral-Obrien et al., 2021). At the same time,
gated, two, three, four, and six subpopulations were revealed, landscape simplification and biotic homogenization have also
respectively. High levels of redundancy were also uncov- altered the nutritional landscape available to bees (Hendrickx
ered. Some core collections had been established because et al., 2007; Lau et al., 2023). With widespread pollinator
none of these studies covered the world collection of sweet- population decline due to interacting stressors, which include
potato germplasm, and DNA samples should be exchanged habitat fragmentation and pesticide use, intentional manage-
and collected to re-evaluate genetic diversity and population ment practices to promote flowering resources in large-scale
structure. The new core collection should be re-established agriculture to improve the overall ecosystem are critical for
using re-evaluation data for future utilization. mitigating the stressors contributing to declining pollina-
Distribution: In theory, sweetpotato germplasm accessions tor health (Crone & Grozinger, 2021; DeGrandi-Hoffman &
should be freely distributed or exchanged nationally and Chen, 2015; Di Pasquale et al., 2013; Dolezal & Toth, 2018).
internationally if the request is eligible. There are some This interaction is an opportunity to promote beneficial land-
restrictions due to the germplasm distribution and exchange scape for both pollinators and farmers to investigate the role
policy between states and countries. Even during the COVID- of sweetpotatoes in the agroecosystem, an attractive resource
19 pandemic, Plant Genetic Resources Conservation Unit for honey bees, bumble bees, and other solitary bees (Wolfe,
distributed many sweetpotato germplasm accessions to differ- 1992). In many cases, sweetpotatoes are planted along with
ent states (Table 2) for breeding programs and genetic studies. other mass cropping systems, including soybeans, corn, and
There are two alarm signals that should always be kept in cotton, and can provide bees with valuable resources in a crit-
mind. One is not to distribute pathogen-infected accessions, ical period of summer resource dearth (Dolezal et al., 2019;
and another is to control the invasive (threaten other plants) Lau et al., 2019).

GEORGE ET AL.
Further research is needed to understand the plant–insect
interactions between sweetpotatoes and bees to determine
if sweetpotatoes can serve as a resource for bee nutrition.
First, it is vital to compare bee diversity and bee health in
locations where sweetpotato plots are present compared to
sites on a similar row crop landscape without sweetpotatoes.
This would provide information on the benefits of intercrop-
ping sweetpotatoes on pollinator communities. We can then
expand on understanding the resource value sweetpotatoes
add to the nutritional landscape for pollinators. Each plant
species can produce pollen and/or nectar with a unique nutri-
tional profile (Lau et al., 2022; Roulston & Cane, 2000). In
many cases, a monofloral diet containing resources from a
single species is suboptimal for bee health, as the diet can
be deficient in a particular nutrient. Studies are required to
understand the nutritional profiles of the pollen and nectar
of sweetpotatoes, and whether sweetpotatoes can supplement
the healthy landscape for pollinators. Also, metabolomics
methods can be used to determine if the sweetpotato pollen
metabolome is altered under drought stress. Since bees use
floral volatiles as olfactory cues to find resources, the flo-
ral volatile organic compounds can be measured by using
solid phase-microextraction (SPME) and analyzed using gas
chromatography–mass spectrometry (GC–MS) (Rering et al.,
2020; Silva et al., 2018).
Possibilities of using honey bees as an environmental
biomonitor for early detection of sweetpotato pathogens
need to be explored. Sweetpotatoes are plagued by vari-
ous pathogens, including fungal diseases, bacterial diseases,
viruses, and phytoplasma diseases (Hedge et al., 2012). Some
of these diseases, such as the fungus causing stem rot, Fusar-
ium oxysporum, have symptoms that are difficult to detect.
Honey bees are the most important managed pollinator and
can also be an informative biomonitor of plant diseases. Pre-
vious studies used front porch pollen traps to biomonitor
pathogens found in sweetpotatoes (Cunningham et al., 2022;
Tremblay et al., 2019). These traps are designed to knock off
and collect the pollen pellet of a returning foraging bee. The
pollen can then be sorted by color and texture to separate
potential pollen collected from other sources and identified
using traditional palynological techniques to morphologically
identify pollen using morphological features and reference
collections (Jones, 2014; Lau et al., 2019). Fresh nectar can
also be collected during the sweetpotato flowering period.
Both nectar and pollen can be analyzed for sweetpotato
pathogens with targeted approaches, such as polymerase
chain reaction (PCR) or enzyme-linked immunosorbent assay
(ELISA) assays. Alternatively, high-throughput sequencing
approaches can be used to detect and discover pathogens in
sweetpotatoes.
Research and management considerations for various
stakeholders need to be accounted. Farmers may not want to
dedicate important land and resources to a cause that may
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9
not generate any profit. Supporting pollinator populations in
a landscape can net positive returns for both beekeepers and
farmers, as bees’ pollination services improve crop yields
in many different crops (Esquivel et al., 2021; Kral-O’Brein
et al., 2021). Including sweetpotatoes in large row-crop sys-
tems can be a holistic approach with benefits to both farmers
and beekeepers.
4 PEST AND DISEASE MANAGEMENT
IN SWEETPOTATO PRODUCTION
4.1 Sweetpotato pests and IPM
US sweetpotato production in 2022 totaled 25.9 million hun-
dredweight (cwt), an 11% reduction from 29.1 million cwt
in 2021 and down 14% from 30.1 million cwt in 2020
(USDA-NASS, 2023). Sweetpotato farming on a smallholder
scale has also diminished over the years in the sweetpotato-
producing states in the southern United States. Higher labor
costs, unavailability of desirable planting materials, and insec-
ticide resistance associated with repeated applications of
insecticides have all contributed to this reduced production.
Furthermore, competition with large-scale production, high
input costs, and low return have discouraged sweetpotato
farming on smallholder scale that is 200 acres or less in pro-
duction. The higher costs of insecticide applications also pose
a further financial burden on resource-limited farmers and
reduce profitability. The effects of climate change, such as
higher temperatures, increased level of CO2 , and prolonged
periods of drought, warrant investigation into the impact of
both biotic and abiotic stress factors on the intensity of insect
pests and diseases of sweetpotatoes and the crop yield (Quiroz
et al., 2018). Research has shown that these biotic and abiotic
stress factors affect the quality and yield of sweetpotatoes and
leave them prone to insect pest and disease attacks (Imbo et al.,
2016).
Several sweetpotato cultivars exist that are resistant to
soil insects (wireworm) and foliar feeding insects (cucum-
ber beetles, flea beetles). Unfortunately, they do not have the
desirable agronomical traits (deep orange flesh color, wide
adaptation, high yield) as the industry standard cultivars,
Beauregard and Covington.
Numerous insect pests, mostly from the order Coleoptera
attack sweetpotatoes in the United States (Cuthbert & Davis,
1970; Jackson & Bohac, 2006c; Jansson et al., 1990; Schalk
et al., 1991; Sorensen, 2009). Most of these insects attack
sweetpotato roots under the soil surface and they are well pro-
tected, rendering it difficult to control them with the available
soil insecticides. Excessive infestations of sweetpotatoes by
the foliar feeding insects indirectly reduce the plant yield, but
can be easily controlled using foliar insecticides. Significant
damage to sweetpotato roots is caused by immature or grub
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10 GEORGE ET AL.

TA B L E 3 Common insect pests of sweetpotatoes in the United States and type of damage they cause to sweetpotatoes.

Insect species Order/family Common name Type of damage

Conoderus vespertinus Coleoptera/Elateridae Tobacco wireworm Storage roots
Conoderus bellus Coleoptera/Elateridae – Storage roots
Conoderus falli Coleoptera/Elateridae Southern potato wireworm Storage roots
Melanotus communis Coleoptera/Elateridae Corn wireworm Storage roots
Phyllophaga ephilida Coleoptera/Scarabaeidae White grub Storage roots
Euetheola humilis Coleoptera/Scarabaeidae Sugarcane beetle Storage roots
Diabrotica balteata Coleoptera/Chrysomelidae Banded cucumber beetle Storage roots/foliage
Diabrotica Coleoptera/Chrysomelidae Spotted cucumber beetle Storage roots/foliage
undecimpunctata
Chaetocnema confinis Coleoptera/Chrysomelidae Flea beetle Storage roots/foliage
Chaetocnema denticulata Coleoptera/Chrysomelidae Toothed flea beetle Storage roots/foliage
Systena frontalis Coleoptera/Chrysomelidae Red-headed flea beetle Storage roots/foliage
Systena elongata Coleoptera/Chrysomelidae Elongate flea beetle Storage roots/foliage
Cylas formicarius Coleoptera/Brentidae Sweetpotato weevil Storage roots
Naupactus leucoloma Coleoptera/Curculionidae Whitefringed beetle Foliage
Bemisia tabaci Hemiptera/Aleyrodidae Sweetpotato whitefly Foliage
Aphis gossypii Hemiptera/Aphididae Cotton aphid Foliage
Spodoptera frugiperda Lepidoptera/Noctuidae Fall armyworm Foliage

stages of several species of soil insects (Table 3) (Reed et al.,
2009; Sorenson, 2009).
Here, we will discuss the management practices for three
major pests that cause significant economic damage to
sweetpotatoes.
role among risk assessment techniques available for sampling
this pest. Identifying a pheromone will provide base infor-
mation on their chemical ecology and a reliable method to
detect when adults move into the area so that the management
practices can be applied more effectively.

4.1.1 Wireworms 4.1.2 Sweetpotato weevil

Wireworms, the larvae of click beetles (Conoderus spp.), are
highly polyphagous with a wide host range consisting of veg-
etables, grains, and other crops such as peanuts, strawberries,
tobacco, and so forth. They are capable of inflicting economic
damage to the roots and tubers due to their long life cycle
and belowground protected habitat. In southeastern US states,
Conoderus spp. are considered major insect pests (Rashid
et al., 2010; Reed et al., 2009; Willis et al., 2010). The esti-
mation of wireworm population by soil core sampling or soil
bait sampling methods is complex and due to their cryptic
life cycle, chemical control efforts are usually unsuccessful
(Parker & Howard, 2001; Toth, 2013). Numerous studies on
chemical ecology of different species of click beetles have
been done in European countries and Japan (Toth, 2013;
Toth et al., 2008, 2014). No current information is available
about the chemical ecology of the Conoderus spp. There-
fore, identifying any semiochemicals from this genus of click
beetles and/or their host plants will provide first-hand infor-
mation on their chemical communication. This can lead to the
development of pheromone traps, which can play an essential
The sweetpotato weevil (Cylas formicarius F.) (Coleoptera:
Brentidae) is considered the most severe pest of sweetpota-
toes in the field and in storage. Throughout their life cycle,
these weevils can cause feeding damage to sweetpotato roots,
stems, and leaves. Females lay eggs by creating holes inside
the sweetpotato roots. Excessive damage to the roots occurs
due to the tunneling larvae, making them unacceptable for
consumption. Even low numbers of larvae reduce sweetpotato
quality and marketable yield. The larval development time
is highly variable ranging from 12 to 154 days; therefore,
even at low densities, the weevil can greatly reduce the mar-
ketable yield of sweetpotatoes (Sutherland, 1986). Pupation
occurs within the sweetpotato and lasts for 5–11 days. The
reproduction process is completed in the stems and the roots.
Sweetpotato weevils are found throughout the southern US
states from North Carolina to Texas and in the tropical regions
worldwide. In Mississippi, the area north of interstate 20 has
been declared weevil-free zone (Mississippi Department of
Agriculture and Commerce, 2022), whereas the southern part
of the state is in sweetpotato weevil quarantine zone. The

GEORGE ET AL.
sweetpotatoes grown in the south cannot be shipped to weevil-
free areas in the northern states, which constitute major
market for sweetpotatoes.
A heavy infestation of weevils can turn the vines yellow-
ish, which is considered as a symptom of weevil infestation
(Jansson & Raman, 1991; Kuriwada et al., 2013). De-vining
at harvest is essential to keep the weevil populations low for
the next season. Sweetpotatoes should never be left in the field
unharvested. Weevils may also survive the winter in the stored
storage roots that will be used as seed stock for the following
season’s planting. Application of Malathion can kill the over-
wintering weevils in seed storage areas. A 5% Imidan dust
application on sweetpotato roots before storage can provide
effective control of weevils (Hammond et al., 2003). Good
agricultural practices should be used such as planting certified
clean seeds (virus and weevil-free), crop rotation, and burning
crop residues.
Synthetic pheromones of sweetpotato weevil [(Z)-3-
dodecen-1-ol (E)-2-butenoate] have been isolated and used
widely for weevil monitoring under field conditions (Jackson
& Bohac, 2006b; Reddy et al., 2012b; Sureda et al., 2006).
Recent studies have also reported that sex pheromone-baited
plastic pole traps caught 60%–78% more weevils than sex
pheromone-baited delta traps, wing traps, or unitraps (Dilip-
kumar et al., 2019). Successful eradication of these weevils
has been reported using a combination of male annihilation
techniques and sterile insect technique in Kume Island, Japan
(Himuro et al., 2022). Use of male annihilation techniques
reduced the wild population density in a few years, followed
by the release of millions of sterile weevils, which helped to
completely eradicate the weevils from Kume Island. Use of
pheromones and other semiochemicals will be highly benefi-
cial in monitoring and management of these weevils in newly
infested locations.
4.1.3 Plant parasitic nematodes
Plant parasitic nematodes (PPNs) are ubiquitous in soils
around the world, infecting nearly every cultivated crop on
earth, and are estimated to cause $157 billion in yield losses
annually (Abad et al., 2008). As a root crop, sweetpotato
is particularly susceptible to quality issues that may result
from PPN infections. Though dozens of PPN genera have
been reported to be associated with sweetpotato, two gen-
era, root-knot nematodes (Meloidogyne spp.) and the reniform
nematode (RN; Rotolyculus reniformis), are the most preva-
lent and yield limiting on sweetpotato in the United States
and many other regions around the world (Karuri et.al., 2017;
Scurrah et al., 2005).
Root knot-nematodes (RKNs) affect a wide range of veg-
etables, row crops, and perennial plants (Nyczepir & Thomas,
2009). Even low levels of RKNs in the soil can reduce crop
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11
F I G U R E 3 Root-knot nematode (RKN, Meloidogyne enterolobii)
infected sweetpotato cultivar Beauregard. In addition to reducing
sweetpotato yield, this type of severe galling caused by high populations
of RKN in the soil can also render storage roots unmarketable.
yield and the quality of roots (Karssen et al., 2013), and RKNs
are found in nearly all areas where sweetpotatoes are grown
(Clark et al., 2013; Stirling et al., 2020). Aboveground symp-
toms of RKN infection are nonspecific: chlorosis, necrosis,
plant stunting, or wilting as the plant is under duress. The
distinguishing symptoms of RKN damage are underground,
where the roots have been infected and galled (Figure 3)
(Karssen et al., 2013). The nematodes invade the plant and
complete their life cycle within the storage root, leading to
misshaped sweetpotatoes with galling and lesions on the sur-
face. In some instances of infection, cracking of the storage
root can also occur (Clark et al., 2013; Quesada-Ocampo,
2018).
Historically, three closely related RKN species have caused
the most yield losses in the tropical and subtropical environ-
ments where sweetpotato is grown: Meloidogyne incognita,
Meloidogyne javanica, and Meloidogyne arenaria. More
recently, a new RKN species, Meloidogyne enterolobii, has
emerged as a severe pest due to its extensive host range,
high level of virulence, and capability to overcome traditional
RKN-resistant genes in a wide array of agricultural crops
(Brito et al., 2020; Philbrick et al., 2020) (Figure 3). First
reported in the continental United States in Florida in 2002,
M. enterolobii has since spread to North Carolina, South Car-
olina, Louisiana, and Georgia (Rutter et al., 2019; Ye et al.,
2013). This nematode is suspected to be spread by the ship-
ment of infected seed roots used to plant crop fields annually
(Quesada-Ocampo, 2018, 2019; Silva et al., 2021). Popular
RKN-resistant sweetpotato cultivars used to manage endemic
species of RKN have been shown to be ineffective against M.
enterolobii (Rutter et al., 2019). The quick spread of this inva-
sive RKN species has prompted significant concerns among
growers and regulators, leading to the imposition of multiple
interstate quarantines on sweetpotato seed roots (FINDMe,
2022; Hare, 2019). Distinguishing between M. enterolobii and

12
the endemic RKN species requires a molecular test conducted
by a trained pathologist, and growers should consult their state
extension agents if they suspect they have a problem with any
RKN.
It can be difficult for a grower to manage RKN, once a field
has become infected. Various factors, including soil compo-
sition, temperature, average rainfall, and host crop rotations,
can influence RKN population densities. RKNs adapt to many
host plants and environments and can continue to reproduce
successfully even as host plants are rotated across crop-
growing seasons (Castagnone-Sereno et al., 2013). Though
rotations to specific nonhost cover crops such as sunn hemp
can significantly reduce RKN populations, they can quickly
rebound once a highly susceptible host is replanted. Some
popular sweetpotato cultivars, such as Covington, offer mod-
erate resistance to the endemic RKN species and can help
keep populations down. However, these same RKN-resistant
cultivars are highly susceptible to M. enterolobii. Multiple
resistant germplasms have been reported in the USDA-GRIN
collection (Rutter et al., 2021; Schwarz et al., 2021). However,
no M. enterolobii-resistant cultivars have yet been released for
the commercial market in the United States.
With minimal management options currently available to
control M. enterolobii in sweetpotato, the best option for
growers is to prevent infection and spread of this nematode in
their fields. The best way to prevent infection by M. enterolo-
bii is to use clean “seed” material for planting that has been
certified by a state agency (FINDMe, 2022; North Carolina
Sweetpotato Commission, 2022). Though certified clean seed
is more expensive, it is grown directly as either first- or
second-generation material out of sterile tissue culture. It has
been grown in fields that are monitored for the presence of M.
enterolobii and other sweetpotato pathogens. Though noncer-
tified roots may look unblemished, they may still harbor low
levels of RKN and other pathogens that can permanently infest
a field.
RN (R. reniformis) is another species of PPN that causes
severe yield and quality issues in sweetpotato. It is the second
most damaging PPN of sweetpotato in the southern United
States (Abel et al., 2007; Smith et al., 2017). RN greatly
reduces storage roots’ quantity, size, and quality upon infec-
tion. Heavily infested sweetpotato storage roots are more
likely to crack, and yield reductions have been noted to
occur without cracking (Smith et al., 2017). Beyond para-
sitic females on the root itself, RN lacks any specific root
symptomology, and foliar symptoms often resemble nutrient
deficiency or water stress (Smith et al., 2017). This lack of
distinctive symptoms has likely led to extensive misdiagnosis
of RN infection.
Management tools for RN in sweetpotato are limited. Crop
rotation is not as effective for this nematode because of its
ability to persist in soil for up to 20 years in the absence of a
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GEORGE ET AL.
suitable host and its ability to quickly rebound to a damaging
level when a susceptible host becomes available (Khanal et al.,
2018; Smith et al., 2017). A few fumigant nematicides can
suppress RN populations, but the added cost of these nemati-
cides and their adverse effects on health and the environment
make them a poor option. The use of host plant resistance
is the most economical approach to managing nematodes in
the field. However, there are currently no RN-resistant sweet-
potato cultivars available in the United States. As with any
nematode problem, growers should sample their fields at the
end of the growing season, preferably right before harvest,
and send them to a state lab for quantification. Nematode
soil counts can provide growers with valuable information on
whether nematicide use is justified in the following growing
season. And just as with RKN, using certified clean seed can
also reduce the possibility of new fields becoming infested
with this nematode.
4.2 Host plant resistance in sweetpotatoes
to ground-dwelling insect pests
Host plant resistance to most sweetpotato insect pests has been
identified (Jackson & Bohac, 2006a, 2006b; Jackson et al.,
2002, 2010, 2012; Wadl et al., 2022). In general, resistance
to insects is not associated with undesirable traits in sweet-
potato (Jones & Cuthbert, 1973), except for a reduction in
final yield (Jackson et al., 2002). A long-term breeding pro-
gram at the USDA-ARS, United States Vegetable Laboratory,
has developed orange-fleshed sweetpotatoes for the primary
domestic market and dry-fleshed sweetpotatoes for use in
value-added products (processing). The program has been
successful in creating high-quality orange-fleshed (Bohac
et al., 2007; Collins et al., 1991; Jackson et al., 2010; Jones
et al., 1986) and dry-fleshed (Bohac et al., 2001; Jackson
et al., 2011) breeding lines and cultivars with resistance to
the common ground-dwelling insect pests of sweetpotato.
Although cultivars and germplasm lines with high levels of
multiple resistance have been released, these insect-resistant
sweetpotato releases have not achieved commercial appeal
due to their low yield, insufficient short maturity, and lack of
disease resistance attributes. Research indicates ample phe-
notypic (Jackson et al., 2018, 2019, 2020; Wadl et al., 2022)
and genotypic (Slonecki et al., 2023; Wadl et al., 2018)
diversity within the USDA sweetpotato germplasm collec-
tions. This diversity, coupled with the existing sources of
insect resistance, provides a foundation for the continuation of
progress in developing improved sweetpotato with resistance
to ground-dwelling insect pests. Furthermore, host plant resis-
tance in sweetpotato offers an environmentally friendly IPM
approach that can reduce the impact of pesticides, which can
be expensive, unreliable, and toxic.

GEORGE ET AL.
4.3 Sweetpotato diseases and virus complex
affecting sweetpotatoes
Sweetpotato is subject to many diseases caused by bacteria,
fungi, and viruses. In general, bacterial diseases do not impact
sweetpotato production. Many fungal diseases can infect the
sweetpotato crop in different developmental stages. Major
diseases on plant bed are Sclerotial blight (Sclerotium rolf-
sii), Slime molds (Fuligo violacea or Physarum plumbeum),
and Rhizoctonia stem canker (Rhizoctonia solani). Many seri-
ous fungal diseases occur in field production of sweetpotato,
including black rot (Ceratocystis fimbriata), foot rot (Plen-
odomus destruens), Fusarium root rot and stem canker and
surface rot (Fusarium oxysporum, Fusarium solani), Fusar-
ium wilt (Fusarium oxysporum f. sp. batatas), and Scurf
(Monilochaetes infuscans). Vast majority of storage rot of
sweetpotato is caused by fungi, including Rhizopus soft rot
(Rhizopus stolonifer), Java black rot (Diplodia gossypina),
dry rot (Diaporthe phaseolorum), punky rot (Trichoderma
koningii), Alternaria rot (Alternaria spp.), blue mold rot
(Penicillium spp.), and gray mold rot (Botrytis cinerea).
The most challenging diseases of sweetpotato are those
caused by viruses due to vegetative propagation. Vegetative
propagation may lead to accumulation of pathogens, partic-
ularly viruses, in the planting stock, resulting in decline in
yield and sometimes quality of the crop (Clark et al., 2012).
The lack of readily available virus-free planting material has
remained a major limiting factor to sweetpotato production
worldwide. Viruses are considered the second most important
limiting factor (after weevils) to sweetpotato production.
Detailed studies have determined the incidence and distri-
bution of sweetpotato viruses in several countries (Abad et al.,
2007; Kwak et al., 2007; Mbewe et al., 2021; Sivaprasad &
Gubba, 2013). According to Clark et al. (2012), the identified
viruses have been assigned to nine families as follows: Bro-
moviridae (1 virus), Bunyaviridae (1), Caulimoviridae (3),
Closteroviridae (1), Comoviridae (1), Flexiviridae (1), Gemi-
niviridae (15), Luteoviridae (1), and Potyviridae (9) (Table 4).
Both aphids and whiteflies transmit majority of the viruses in
sweetpotato. Several studies have confirmed that weed species
play a role in the epidemiology of some sweetpotato viruses
(Akel et al., 2010; Tugume et al., 2010).
A common virus is the aphid-transmitted sweetpotato
feathery mottle virus (SPFMV; Potyvirus, Potyviridae)
(Syller, 2014). SPFMV causes transient symptoms when
infecting alone; it is most damaging in mixed infections when
it is synergized by co-infection with whitefly transmitted,
phloem-limited sweetpotato chlorotic stunt virus (SPCSV)
(Gibson & Kreuze, 2015). The synergistic interaction gener-
ally causes severe “sweetpotato virus disease” (SPVD; Gubba
& Sivaprasad, 2015; Byamukama et al., 2004; Gibson et al.,
1998), which is considered the most devastating viral dis-
ease worldwide. However, other virus combinations may
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13
cause symptoms that resemble SPVD. Depending on where
sweetpotato is grown, different virus complexes have been
identified to infect the crop (Clark et al., 2012). In temper-
ate regions, the crop is generally affected by a complex of
potyviruses and possibly other unknown viruses that typi-
cally cause yield reductions of about 20%–40% (Clark & Hoy,
2006; Clark et al., 2010). In East Africa, SPVD can cause
80%–90% losses in many high-yielding genotypes (Karyeija
et al., 1998). SPCSV, on the other hand, is the most damag-
ing virus causing permanent symptoms even when infecting
alone. In some cases, SPVD can cause yield reductions of up
to 98% (Mukasa et al., 2003).
With the adoption of molecular methods for virus research
in the last two decades, there is a better understanding
on the composition of sweetpotato virus complexes, the
effects of virus diseases on production systems, the biol-
ogy of the virus–plant interaction, and the management
approaches of viral diseases. Since the initial identifica-
tion of the sweetpotato leaf curl virus (SPLCV) (Lotrakul
& Valverde, 1999; Lotrakul et al., 1998), a high genetic
diversity of various begomoviruses has been identified on
sweetpotato (Zhang & Ling, 2011). These emerging bego-
moviruses have caused severe yield losses of up to 60%
compared to those of virus-free materials (Ling et al., 2010).
With no virus-resistant cultivars available, planting virus-
free materials is crucial to ensure sweetpotato production.
Due to the importance of numerous viral disease prob-
lems, sweetpotato foundation “clean seed” programs were
developed long before the advent of technology to produce
virus-free propagating materials. In 2015, the National Clean
Plant Network (NCPN) for sweetpotato was established under
the umbrella of the United States Department of Agricul-
ture (https://www.aphis.usda.gov/aphis/ourfocus/planthealth/
ppa-ppdmdpp/sa_ncpn). Currently, sweetpotato clean seed
programs have been established in California, Louisiana,
North Carolina, Mississippi, Arkansas, and Hawaii (https://
ucanr.edu/sites/ncpnsweetpotato). They apply the meristem
shoot tip culture technique to generate virus-free materials for
local cultivars (Alconero et al., 1975).
Meristem-tip culture starts with the excision of the shoot’s
organized apex from a selected donor plant for subsequent
in vitro culture. The excised meristem tip is typically small
(often <1 mm in length), which holds the potential to exclude
pathogenic organisms that may have been present in the
donor plants. Thermotherapy is commonly carried out at
intervals of every 6 h, alternating between temperatures of
31 and 36˚C, for 3–4 weeks before meristem-tip culture.
This process is repeated and applied to the tissue-cultured
plantlets to partially deactivate the viruses and slow down
their movement. The ability to produce and maintain plants
free of detectable viruses through meristem-tip culture has
dramatically improved sweetpotato yields for several decades.
However, as viruses can accumulate and transfer from one
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14 GEORGE ET AL.

TA B L E 4 Important sweetpotato viruses and their mode of transmission.

Family Genus Virus species Transmission/vector

Bromoviridae Cucumovirus Cucumber mosaic virus Aphid
Bunyaviridae Phlebovirus Sweetpotato C-3 virus Unknown
Caulimoviridae Badnavirus Sweetpotato pakakuy virus Unknown
Closteroviridae Crinivirus Sweetpotato chlorotic stunt virus Whitefly
Comoviridae Nepovirus Sweetpotato ringspot virus Nematode
Flexiviridae Carlavirus Sweetpotato chlorotic fleck virus Unknown
Geminiviridae Begomovirus Sweetpotato leaf curl virus and 14 other sweetpotato viruses Whitefly
Luteoviridae Polerovirus Sweetpotato leaf speckling virus Aphid
Potyviridae Potyvirus Sweetpotato feathery mottle virus; sweetpotato virus C; sweetpotato virus 2; Aphid
sweetpotato virus G
Ipomovirus Sweetpotato mild mottle virus; sweetpotato yellow dwarf virus Unknown

generation to the next through infected vegetatively prop-
agated materials (also known as “seeds”), studies indicate
that a significant reduction of yield and quality may occur
due to re-introduction of viruses from previous propagating
materials. In two separate studies, 100% of virus-indexed
plants were re-infected by SPFMV within the first year in the
field, and a decline in yield occurred gradually over several
years (Bryan et al., 2003). Therefore, sweetpotato “seeds,”
the primary method to reduce the damage of virus infections,
are very expensive because farmers must regularly purchase
virus-tested “seeds” due to the high re-infection rates in the
field.
Factors such as virus variation, time, and required expendi-
ture have mired conventional breeding efforts (Lomonossoff,
1995). Moreover, genetic sources of resistance are scarce and
the incorporation of such resistance from the wild diploid
Ipomoea spp. species into polyploid sweetpotato is a compli-
cated task. With the development of genetic transformation
systems, genome sequences, and genetic engineering tools,
for example, CRISPR (Clustered Regularly Interspaced Short
Palindromic Repeats)/Cas9, molecular breeding provides a
promising strategy for the development of novel cultivars
with value-added traits. The CRISPR/Cas9 system works to
induce targeted genetic modifications to regulate endoge-
nous gene expression, and the “transgene-free” end products
set it apart from traditional genetically modified organisms
(GMO) where foreign gene(s) are integrated into the host
genome (Chaudhary et al., 2018). The CRISPR/Cas9 sys-
tem has ushered in the beginning of a new era in basic
and applied biological sciences (Bomgardner, 2017; Reis
et al., 2014). By using the CRISPR-Cas13 technique, a recent
study demonstrated sweetpotato lines with enhanced viral
disease (SPVD) resistance by targeting one of its essential
pathogeneses-related factors (i.e., SPCSV-RNase3) (Yu et
al., 2022).
The solution to the problems caused by sweetpotato viruses
is to ensure that growers plant virus-indexed clean propaga-
tion material or “seeds.” However, by estimate, there is a 72%
shortfall of clean plant units needed to cover all sweetpotato
acreage (NCPN Network News, May 2018). Thus, there is a
considerable demand for establishing certified nursery farms
to produce clean “seeds” in the sweetpotato industry. Besides,
the high reinfection rate of viruses in production requires
the farmers to frequently purchase virus-indexed propaga-
tion material, which significantly increases their financial
input for propagation and production. Therefore, using other
biotechnological approaches, such as gene transformation or
genome editing techniques, to produce virus-resistant culti-
vars would be another promising strategy to control the virus
in sweetpotato.
5 WEED MANAGEMENT IN
SWEETPOTATO PRODUCTION
Weed management is consistently ranked among the top
research priorities of the US sweetpotato industry. Amaran-
thus species can reduce yields up to 85% in sweetpotato
(Basinger et al., 2019; Smith et al., 2020). Yellow and
purple nutsedge (Cyperus esculentus L. and Cyperus rotun-
dus L., respectively, Family: Cyperaceae) negatively affect
sweetpotato yield and quality, and losses from 18% to 96%
have been reported (Meyers & Shankle, 2015). Large crab-
grass [Digitaria sanguinalis (L.) Scop.] at densities of 1–16
plants·m−1 of row reduced yields from 35% to 76% in sweet-
potato (Basinger et al., 2019). Grieg and Al-Tikriti (1966)
and Glaze et al. (1981) found that yields of sweetpotato plots
were reduced by over 90% in comparison with treatment plots
receiving herbicides, hand weeding, and cultivation. Glaze
et al. (1981) reported that Georgia Red sweetpotato yields

GEORGE ET AL.
were reduced by 90% when weedy control plots were com-
pared to plots receiving cultivation and herbicides in 1 year of
the study.
Conventional sweetpotato growers utilize herbicides,
between-row cultivation, mowing, and hand removal to
manage weeds. Currently, 10 herbicides are registered for
weed control. Those commonly used are flumioxazin, S-
metolachlor, clomazone, and two graminicides (sethoxydim
and clethodim). Although napropamide and DCPA are reg-
istered for sweetpotato, they provide inconsistent and often
inadequate weed control (Weir, 2001). Each of the registered
herbicides has drawbacks. Flumioxazin, S-metolachlor, and
clomazone require rainfall or irrigation for activation, but
few producers have the infrastructure for overhead irrigation.
If rainfall is not timely, weeds emerge before activation
and are not controlled. Flumioxazin must be applied before
transplanting and requires that planting ridges be formed
and then the top of the ridge leveled. If not done correctly,
the herbicide is removed from the center of the planted row
during transplanting, thereby providing little weed control.
Weeds that escape control in the row cannot be controlled
with cultivation and compete with the developing crop.
In organic and/or resource-limited production systems,
weed management is more difficult because the use of syn-
thetic herbicides is prohibited or too expensive. Mechanical
weed control is a common practice among these sweetpotato
producers, who disc multiple times during field preparation
and cultivate two to three times during the growing season.
Escaped weeds are removed by hand. Many organic fields in
the Southeastern United States are hand weeded at an esti-
mated expense of $510 per acre. The lack of adequate weed
control is the most critical obstacle to adopting organic pro-
duction or sustainable cultural practices (i.e., no tillage or
minimum tillage). Crop rotation to manage weeds can be suc-
cessful if the crops are more competitive with the target weed
species and a minimum of 2 years is often required to sig-
nificantly reduce weed pressure before planting sweetpotato
again (Monks et al., 2018).
Cultivars tolerant to weed interference can be essential
components in integrated weed management in both conven-
tional and organic production. The leading US sweetpotato
cultivars (Beauregard and Covington) are highly suscepti-
ble to weed interference to the extent that total crop failure
has been reported (Meyers et al., 2010). Delaying weeding
beyond 2 weeks after planting resulted in substantial reduc-
tion in yield (Levett, 1992; Seem et al., 2003). Yields of
Carolina Bunch, a cultivar with semi-erect vine growth habit
(Huamán, 1991), were reduced by ≤20% by weed interference
in comparison to weed-free plots, whereas all other clones
were reduced from 50% and 70% (La Bonte et al., 1999).
Sweetpotatoes with erect to semi-erect growth have shorter
internodes, which results in a denser canopy with greater
height and more significant branching in the early growth
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15
stages. These clones with erect to semi-erect plant habit grow
radially and form a closed canopy earlier than cultivars with
a prostrate spreading plant habit, thus improving weed sup-
pression. Harrison and Jackson (2011) compared Carolina
Bunch and Beauregard (spreading habit) in weed-free plots
and reported yield reduction in Beauregard. Development
of erect and semi-erect vined sweetpotato germplasm could
allow increased tillage later into the season as the vines
would not wholly cover between the rows. To provide addi-
tional weed and insect management strategies for sweetpotato,
research is required for the development of insect-resistant
germplasm that also has competitive weed tolerance poten-
tial by breeding and selecting for sweetpotato clones that are
fast growing and have semi-erect to erect canopy architecture.
Research studies have reported that some cultivars may be
more tolerant to weeds than others (Jackson et al., 2011; La
Bonte et al., 1999). A study at United States Vegetable Labo-
ratory compared the performance of six advanced sweetpotato
clones to three control cultivars (Beauregard, Covington, and
Monaco) over two seasons under various weed-free intervals
and highlight the potential for the development of germplasm
with tolerance to weed interference and resistance to insect
pests (Wadl et al., 2022). Additional research is needed to
address the need for multi-year field trials for evaluating insect
resistance, lack of existing germplasm with erect plant habit,
lack of sufficient weed pressure each year, and impact of
abiotic stress (drought and flood) on yield.
6 WATER MANAGEMENT IN
SWEETPOTATO PRODUCTION
Sweetpotato is considered a drought-tolerant crop (Khan &
Doty, 2009) but also sensitive to water logging (Gomes &
Carr, 2003a). As such, water plays a vital role in its growth
and yield. Little is known about the water requirements and
yield responses of this crop to irrigation. Norman and Molales
(1984) and Gomes and Carr (2003a) commented on how
few studies there have been on the water use (WU) of this
crop despite its many cultivars. Most of the water research
in sweetpotato has been on the effects of water deficit as
most of this crop is produced in areas that are primarily
rain fed, such as the drought-prone tropics. Some cultivars
have been shown to be susceptible to drought (Adebola &
Abe, 2013; Karakas et al., 2021). Water deficits reduce leaf
water potential and total WU, and subsequently reduce stom-
atal conductance, leaf area, root mass, total plant mass, and
tuber yield (Sivan et al., 1996). Van Heerden and Laurie
(2008) examined the effects of long-term restricted water sup-
ply on shoot development, photosynthesis, and storage root
yield of two cultivars in rainout shelter conditions in South
Africa and found that significant decreases in stomatal con-
ductance occurred in both cultivars after 5 weeks of treatment.

16
However, continued measurements revealed a significant cul-
tivar difference in the persistence of this response and its
effects on CO2 assimilation (Van Heerden & Laurie, 2008).
Ekanayake and Collins (2004) found that drought stress sig-
nificantly reduced nitrogenous compounds and root yield
of sweetpotato in Peru. In Thailand, Yooyongwech et al.
(2017) were able to elevate water-deficit tolerance by using a
foliar application of paclobutrazol that improved soluble sugar
and free proline accumulation, photosynthetic pigment sta-
bilization, photosynthetic abilities, growth performance, and
storage root yield.
Several studies have shown contradicting information about
the necessity of irrigation in sweetpotato. Smittle et al. (1990)
showed that marketable yield and yield of US No.1 grade roots
generally decreased when soil water tensions exceeded 25 kPa
before irrigation, while soil water stress of 100 kPa during
storage root development did not significantly affect yield in
a field study in Georgia. Lana and Peterson (1956) showed
similar results with peak yield when irrigated at 50% available
soil moisture. However, in other studies (Bowers et al., 1956;
Ghuman & Lal, 1983), the yield was increased in irrigation
plots compared with nonirrigated plots, but there was no dif-
ference among the different irrigation frequency and quantity
levels examined. Hernandez (1965) also recommended that
the amount of water needed to produce a good crop varies
yearly and that irrigation should commence while the soil
moisture in the root zone is higher than 25%, possibly 40%–
50%. It is often anecdotally recommended that irrigation is
only needed in the early planting stages (first 30 days), though
there is little information available in the literature to indicate
ideal timings and methods of irrigation.
Very little research has been done on WU (the amount
of water used to produce crops in the field) and water-use
efficiency (WUE; the amount of water used in relation to
yield) in sweetpotato, and field studies are minimal in general,
especially in the United States. Kelm et al. (2001) conducted
pot-scale research on the effect of nitrogen deficiency on the
WUE of one Peru cultivar. They showed WUE was dependent
on leaf inclination and chlorophyll content in leaves, which,
in turn, was dependent on nitrogen supply. In 2014, Masango
(2014) published a master’s thesis on WUE in orange-fleshed
sweetpotato in South Africa under rain shelters with different
irrigation regimes and stated WU ranged from 298 to 478 mm.
WUE ranged from 64.8 to 97.5 kg·ha−1 ·mm−1 , while Afzal
et al. (2021) put WU at 500 mm on average worldwide. Gomes
and Carr (2003a) measured WU from well-watered crops as
800 mm during the rains and 550 mm during the dry season,
while the WU for the rain-fed crops was 360 and 180 mm,
respectively. Karakas et al. (2021) found that seasonal water
consumption of two sweetpotato cultivars was calculated as
808 and 826 mm, under no water deficit in Turkey. Gomes
and Carr (2003b) recorded WUEs of 13 kg·ha−1 ·mm−1 in the
rains and 24 kg·ha−1 ·mm−1 in the dry season, indicating each
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GEORGE ET AL.
unit of water “lost” in the dry season was nearly twice as pro-
ductive as the same amount of water consumed by the crop
in the rains. Dladla et al. (2018) examined the effect of ridge
type and environmental conditions on WUE in South Africa
and found that WUE was higher under peaked ridges. They
also stated that cultivars performed differently at each site and
under the different ridge types and suggested producers adopt
different cultivars across different environments to improve
yield and WUE. No numbers for sweetpotato WU or WUE
were found for the United States from the last 30 years.
7 IMPACT OF CLIMATE CHANGE ON
GLOBAL SWEETPOTATO PRODUCTION
Global food security in the 21st century is severely threatened
by climate change and may impact various agricultural pro-
duction systems. It is estimated that climate change will have
positive and negative impacts on agricultural systems glob-
ally, with adverse effects outweighing the positive ones (Bage,
2007). Climate shifts lengthen growing seasons, and rise in
temperatures may bring along negative implications such as
reduced precipitation, thus affecting water availability and,
in turn, crop water requirements (Eitzinger & Kubu, 2009;
Molua & Lambi, 2006). An increase in atmospheric tempera-
ture and elevation of CO2 concentration could influence crop
yield and may directly or indirectly impact crop pests, weeds,
and diseases. Climate variability and change may have a more
significant impact on tropical agricultural production systems
where the temperature is projected to increase from 1.5˚C in
the next 20 years to 4.3˚C by 2080 (Hepworth & Goulden,
2008), leading to changes in the distribution of agroecological
zones, soil moisture, and shortened growing seasons (Hulme,
1996).
7.1 Impact on pest and disease management
Climate change can impact pest and disease occurrences,
host–pathogen interactions, ecology and distribution of
insects, time of appearance, natural enemy populations, insect
migration, and overwintering capacity, becoming a major set-
back to agricultural production (Ayyogari et al., 2014). As
insects are poikilothermic organisms, the increase in atmo-
spheric temperature may directly impact insect behavior,
developmental biology, host selection, reproduction, popula-
tion dynamics, and dispersal mechanisms. Indirectly, climate
change may affect the plant–pest–natural enemy interactions
and their relationships with other insect species, natural ene-
mies, symbionts, and mutualists (Abdallah et al., 2014). These
climatic factors create new ecological niches for insects to
establish and spread in new geographical areas (FAO, 2019).
Studies have reported how temperature changes affect the

GEORGE ET AL.
growth of pest populations in important grain crops such
as wheat, rice, and maize. Deutsch et al. (2018) reported
that global warming would accelerate the growth of pest
populations in wheat grown under temperate conditions
and decrease the growth of pest populations in rice grown
in tropical zones. Maize grown in temperate and tropical
regions may experience a mixed growth response of pests to
global-warming-associated changes. Bale et al. (2002) pre-
dicted that aboveground insects would be more affected by
increased temperatures than belowground insects. As sweet-
potato roots are grown belowground, many root-damaging
pests are belowground and may not be significantly affected
by this temperature increase.
7.2 Impact of CO2 levels on sweetpotato
production and yield
It is estimated that the CO2 concentrations will increase from
400 to >700 ppm by the end of the century (Fahad et al.,
2017). Elevated CO2 has been shown to be beneficial for
C3 plants such as potato, sweetpotato, and yam compared
to C4 plants such as maize and sorghum whose yields espe-
cially for maize were significantly reduced (Raymundo et al.,
2014). The effects of increasing CO2 levels on insect pests
are highly dependent on their host plants (Coviella & Trum-
ble, 1999). These differential effects of elevated atmospheric
CO2 on C3 and C4 plants may result in asymmetric effects on
herbivory, and the response of insects feeding on C4 plants
may differ from that of C3 plants. C3 plants are likely to be
positively affected by elevated CO2 and negatively affected by
insect response, whereas C4 plants are less responsive to ele-
vated CO2 and, therefore, less likely to be affected by changes
in insect feeding behavior (Lincoln et al., 1986; Skendzic
et al., 2021). Studies by Finnan et al. (2005) showed that
potato yields increase with elevated CO2 in open-top cham-
bers (OTCs) and free air carbon dioxide enrichment (FACE)
systems across Europe and the United States. Other stud-
ies indicated that increased temperature may counteract the
positive effect of elevated CO2 in potatoes (Schapendonk
et al., 1995). Biswas et al. (1996) showed that sweetpota-
toes under elevated CO2 in combination with water stress
did not respond to elevated CO2 , while the yield of well-
watered plants increased significantly with elevated CO2 .
This shows that multiple factors and interactions may play a
role in determining yield parameters under varying climatic
factors.
7.3 Impact of precipitation patterns on
sweetpotato production
Changes in precipitation patterns can also influence pest
occurrence and development. Insect species that overwin-
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17
ter in the soil can be directly affected by heavy rainfall and
flooding conditions (Skendzic et al., 2021). Wireworms are a
damaging pest of storage root crops such as potatoes, sweet-
potatoes, and sugar beets that are grown belowground. Staley
et al. (2007) found rapid growth of wireworm populations
in the upper part of the soil due to increased summer rain-
fall events as opposed to ambient and drought conditions.
Yihdego et al. (2019) reported that drought-stressed plants
are more susceptible to insect attack because of decrease
in plant secondary metabolite production that contributes to
plant defense responses.
Because tuber and root crops are a significant food source
and a staple for many developing countries in Africa, many
cropping models predicting climate change’s impact on water
and yield estimates for sweetpotatoes are studied and reported
for sub-Saharan countries. Tuber crops are C3 plants, and their
photosynthesis relies mainly on CO2 concentration (Flexas
& Medrano, 2002). If the crops’ water needs are not met,
the water deficit may lead to stomatal closure, thus reducing
the amount of water lost through evapotranspiration (Blum,
2009). If the soil and plant water status are not replenished
through irrigation, stomatal closure leads to reduced CO2
uptake, which results in reduced biomass production. Under
global warming conditions, an expected increase in tempera-
ture, evapotranspiration, and CO2 may lead to a decrease in
soil moisture affecting the soil–plant water relations (Kimball
& Bernacchi, 2006). Under drought-like conditions, transpira-
tion efficiency is vital in maximizing biomass production and
the crops’ primary productivity through increased CO2 fixa-
tion (Gherardi & Sala, 2020). If these irrigation requirements
during the root bulking stage (mid) of sweetpotatoes are not
met, it may lead to reduced growth and development of tubers
(Kassam & Smith, 2001).
While studies have combined climate, crop, and eco-
nomic models to examine the impact of climate change
on agricultural production and food security, results have
varied widely due to differences in models, scenarios, and
data used in these studies. Understanding the magnitude
of the impacts of climatic factors on sweetpotato produc-
tion is further complicated by the interaction of numerous
biophysical and socioeconomic factors (Raymundo et al.,
2014). Two crop simulation models, MADHURAM (Soma-
sundaram & Santhosh Mithra, 2008) and SPOTCOMS,
which is a MADHURAM model with a modified canopy
algorithm (Santhosh Mithra & Somasundaram, 2008), have
been reported for sweetpotatoes (Raymundo et al., 2014).
The MADHURAM model simulates photosynthesis across
canopy layers to calculate direct and diffused sunlight inter-
ception. This model considers three phenological stages
(initial–middle–final), crop growth, and yield by consider-
ing water, potassium, and nitrogen limitations. Compared to
MADHURAM, SPOTCOMS is a simpler model, although the
canopy development includes branching (Santhosh Mithra &

18
Somasundaram, 2008). In both MADHURAM and SPOT-
COMS, the phenology stages are determined by growing
degree days, with a base temperature of 8˚C, an optimum
temperature of 25˚C, and a maximum temperature of 38˚C.
These values for base temperature appear low as sweetpotato
is cropped in subtropical and tropical regions. No published
studies have reported applications of these two sweetpotato
models, except for Villordon et al. (2009), who used these
models to simulate the harvest dates for sweetpotatoes in
Louisiana.
Using CROPWAT 8.0 model, Mbayaki and Karuku (2021)
predicted the implications of climate change on crop water
requirements for the short rain seasons between 1991 and
2016 (baseline climate) and the future from 2020 to 2039
(climate change) in sweetpotato growing regions of Kenya.
Based on the models, this study predicted that the average
annual temperatures would rise by 36.3% and could shorten
the sweetpotato growth periods by 42 days lowering the yield.
Irrigation water requirements will be increased as the annual
rainfall is supposed to reduce by 16.7%, which may impact
soil moisture and reduce water availability. While Mbayaki
and Karuku (2021) predicted a decrease in rainfall and a
reduction in yield unless the irrigation requirements are not
sufficiently met, Ddumba (2018) indicated a 50-mm increase
in rainfall for most parts of Kenya and Tanzania, and a
decrease in rainfall for central Uganda and regions of south-
ern Tanzania using another SPOTCOM crop model. Based
on these projections, a rise in rainfall will increase sweet-
potato yield by 4 t·ha−1 in western and southern Kenya in
2050 and an overall increase by 1–3 t·ha−1 across the east
African region. This model projected increased precipita-
tion and temperature; therefore, higher sweetpotato yields
of 7, 10, and >20 t·ha−1 were projected for the 2030s,
2050s, and 2070s for four cultivars grown in the east African
regions. Though some of these models have contradicting
findings on precipitation and yield projections of sweetpota-
toes, these studies agree that the atmospheric temperature
is going to rise globally and will have an impact on crop
production.
Crop models can be used effectively to assess the impacts
of climate change and potential adaptations if the models are
well-tested and proven to reproduce the results from field-
based experiments, including variations in climate change
factors (Raymundo et al., 2014). Such crop models are lacking
in tuber crops, and there is an urgent need to develop models
that combine physiological studies on high temperature, heat
stress, CO2 , and water stress, evaluating different cultivars
and their impact on crop growth dynamics, pest and diseases,
nutrient and water uptake, and yield. Collecting such detailed
data on crop growth under varying field conditions and farm-
ing situations will help to develop better agronomic decisions
locally, regionally, and globally.
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GEORGE ET AL.
8 CURRENT LIMITATIONS AND
CHALLENGES FOR SMALL-SCALE AND
ORGANIC FARMERS
Currently, there is no detailed published information on the
demographics, acreage, production practice problems, and
challenges of underserved sweetpotato growers, especially
African American growers. In this regard, the 1890 Small
Farm Working Group is working to address the knowledge
gap and to increase the adoption of new, practical, and
economical IPM practices by underserved small farmers.
The 1890 Small Farm IPM Working Group has informally
surveyed and engaged with over 60 underserved farmers
concerning priorities and challenges, such as climate-smart
IPM and organic production practices via letters of sup-
port, personal engagement, and phone interviews. The main
concerns for organic sweetpotato producers were identified
as (1) labor cost required for organic production, (2) the
efficacy of biopesticides in managing sweetpotato pests,
and financial benefits, (3) limitations on having enough
land for crop rotations or suitable rotational cash crop that
will return an economic benefit, if not growing sweetpota-
toes, and (4) input cost of organic production and markets
for selling organic sweetpotatoes. Also, small and organic
growers face challenges with inadequate number of mar-
ketplaces and proper agricultural infrastructures (i.e., fruit
and vegetable processing facilities). Smallholder growers
also require more technical assistance through research,
extension, and educational programs related to organic pest
management, water and nutrient management, and climate-
smart agricultural practices. Financial management is another
major limitation that limits access of smallholder farm-
ers to capital, land, technology, labor, and experiential
knowledge. There is a strong interest among underserved
farmers to participate in and adopt organic production prac-
tices, developing markets and value-added products for
sweetpotatoes.
8.1 Small farm pest management challenges
Surveys on small-farm IPM challenges are scarce. Pinero and
Keay (2018) conducted an online survey aimed at charac-
terizing farming practices and challenges, methods of pest
management, level of IPM knowledge, and preferred sources
of IPM information of commercial fruit and vegetable pro-
ducers in Missouri. This survey elicited responses from
growers farming in 44 counties across the state and provided
a comprehensive perspective on the scale and scope of pro-
duction of fruit and vegetable crops. When asked to select
the most significant challenge on their farm from among
eight options, 43% of respondents selected plant pests (which

GEORGE ET AL.
comprised diseases, arthropods, and weeds) followed by
weather (21%). Both conventional and organic producers
chose insects as the most important pest of tree-fruit pests,
followed by diseases. In vegetables, insects were chosen
as the most significant problem, followed by weed man-
agement. Both groups selected diagnosing diseases as their
most-significant challenge; however, conventional growers
had more trouble with deciding which fungicide chemistries
to use more often than developing a prevention plan. Over
56% of the farmers had concerns about disease management
for transplants in greenhouses and nurseries. Both conven-
tional (68%) and organic (70%) producers selected insects
as a significant pest of vegetables. Both groups consider
themselves as having a medium knowledge of IPM, possi-
bly reflecting the need or requirements of certified organic
growers to understand and implement plant health manage-
ment strategies such as crop rotations, cover crops, sanitation,
cultural practices, and biological controls as indicated by the
USDA’s National Organic Program final rule (USDA, 2000).
The authors of the small-farm IPM study in Missouri
pointed out that their survey of small farmers was not ethni-
cally diverse, with 94% of respondents describing themselves
as White. This was followed by growers of Asian/Pacific
Islander origin (8%) and Black or African American (3%).
According to the 2012 Census of Agriculture (USDA-NASS,
2014), 99.4% of farms (all farm types) in Missouri were oper-
ated by Whites. In contrast, Hispanics operate only 0.8% of
farms, and 0.28% were operated by either Black or African
American growers. There is a need to survey and document
IPM challenges and farming practices of diverse under-
served small farmers. In other southern states, Black farmers
constitute a larger share of total farmers, with Mississippi
(12%) leading, followed by Louisiana (7%), South Car-
olina (7%), Alabama (6%), and Georgia (4%) (USDA-NASS,
2014). Also, more information is needed on pest manage-
ment challenges and concerns of underserved smallholder
farmers.
8.2 Approaches and solutions for organic
and underserved farmers
Specialty crop growers should adopt climate-smart practices
such as no till, cover crops, biodiversity, and use of biologicals
for establishing consumer-driven markets for climate-smart
commodities. Alcorn State University (Lorman, Mississippi)
is a collaborator on a research and educational project to
facilitate the implementation of climate-smart production
practices by underserved small-farm specialty crop grow-
ers (including sweetpotatoes). This project aims to develop
robust tools for measuring inputs, outputs (climate-smart
ecosystem services), and crop yields and to establish systems
for traceability, verification, and marketing of climate-smart
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19
commodities. Growing sweetpotatoes using climate-smart
agricultural production practices combined with marketing is
a solution to help small, underserved farmers get premium
prices for their produce.
In recent years, organic produce is in high demand and there
is a long history and interest of African American farmers
growing crops organically. However, the transition to organic
production can be a lengthy and expensive process for many
Black farmers. In the 2012 Census of Agriculture (USDA-
NASS, 2014), out of the over 33,000 principal Black farmers,
around 116 were certified organic farmers, which is less than
0.1% of total certified organic farmers (14,093) in the nation.
Mississippi has the highest percentage of African American
specialty crop farmers in the United States. Mississippi, and
other states in the Southeast with significant numbers of Black
specialty crop farmers (LA, AL, SC, and GA), is an important
geographical area to focus on expanding organic sweetpotato
production (Grist, 2016).
9 CHALLENGES AND SUSTAINABLE
APPROACHES FOR IMPROVING
SWEETPOTATO PRODUCTION
Root/tuber crops such as sweetpotatoes have contributed sig-
nificantly to reducing poverty in the developing world and
have enhanced food security by addressing hunger, malnu-
trition, and micronutrient deficiency, thanks to their higher
nutrient content (Afzal et al., 2021). In addition to being a
staple food crop and animal feed, sweetpotatoes have poten-
tial for biofuel production due to its high starch content and
bioethanol yield (Ziska et al., 2009). Sweetpotato cultivation
also generates sustainable income for smallholder farmers
and contributes to the livelihood of farmers due to lower
production costs. Diversification of sweetpotato through pro-
cessed products and value addition can generate extra income
and increase crop utilization. Also, sweetpotato can increase
resilience and reduce the vulnerability of smallholder agricul-
tural production systems to climate change effects and other
disruptions (Afzal et al., 2021). There is a wide genetic diver-
sity of this crop that is maintained in gene banks for farmers
(Anglin et al., 2021; Elameen et al., 2008; Lee et al., 2019;
Slonecki et al., 2023; Su et al., 2017; Wadl et al., 2018).
All these attributes associated with sweetpotato production
make it a sustainable crop for agricultural production systems.
However, there are many challenges related to sweetpotato
production, such as climatic factors, pest and disease prob-
lems, value addition, nutritional quality, marketability, and
cultivar selection that make this crop less attractive to some
growers. Sustainable approaches need to be developed in dif-
ferent areas to attract more growers into this crop and increase
the acreage, production, and value addition of this crop in the
United States and globally.

20
9.1 Exploring new germplasm collections,
sweetpotato chemistries, and their health
benefits
Consumer demand for purple sweetpotatoes has steadily
increased over the last decade. There is more interest from
growers for different cultivars, especially purple sweetpotato
cultivars, but more research is needed for the development of
cultivars and their utilization. Germplasm collections of dif-
ferent cultivars need to be re-evaluated for genetic diversity
and population structure. To expand the genetic diversity of
the collection, newly developed cultivars and advanced breed-
ing lines need to be acquired, evaluated, and characterized for
nutritional quality. Single-nucleotide polymorphism arrays
can be effectively used for the evaluation of germplasm, and
cryopreservation techniques can be employed for germplasm
preservation and overcoming germplasm deterioration. Fur-
ther analytical studies are required to study the effects of
anthocyanin types and contents of purple sweetpotato culti-
vars and to identify compounds that have immunomodulatory
effects. Technological advances in analytical chemistry are
available to characterize the specific molecules in sweetpota-
toes that contribute to specific health-promoting properties
as well as sensory traits. For example, anthocyanin types
and quantities can be characterized using liquid chromatog-
raphy with photo diode array and triple quadrupole mass
spectrometer detectors (LC–PDA–TQMS), and aroma com-
pounds present in the baked roots can be characterized using
comprehensive two-dimensional gas chromatography with a
high-resolution mass spectrometer (GC×GC–ToFMS) and
sensory analysis. Fractionation and purification of these com-
pounds enable further testing in model cell lines to screen
for compounds that are potentially health promoting. These
approaches will help to identify favorable purple sweetpotato
anthocyanins that have health-promoting properties and are
beneficial or inconsequential to the sensory experience. Con-
trarily, anthocyanins that have minimal health benefits and/or
are associated with undesirable sensory attributes will be tar-
geted for reduction in purple sweetpotato breeding selections
and production practices. This will also benefit health sci-
ences, food sciences, and horticulture programs involved with
anthocyanin-containing food products or crops that would like
to decrease anthocyanins with undesirable tastes and increase
those with beneficial attributes (e.g., natural color extracts,
blueberry breeders).
Sweetpotatoes have great potential for value addition
through food processing. However, processing sweetpota-
toes has additional hurdles due to its unique composition
and labile phytonutrients. Some of the US sweetpotato cul-
tivars have a higher sugar content. Therefore, achieving a
crispy texture in fried products (e.g., chips and fries) without
excessive browning and acrylamide production is a challenge.
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GEORGE ET AL.
Identification of sweetpotato attributes (e.g., chemical com-
position, starch attributes, and cell wall structures) that impact
fried sweetpotato textures will aid breeders and processors in
selecting cultivars and how to make the products that meet
consumer preferences. Further research is needed to fully
understand the impact of sweetpotato chemistry on prod-
uct quality and enable the development and optimization of
sweetpotato processing technologies. Improving our knowl-
edge on sweetpotato chemistry and processing research will
bring more value to the sweetpotato industry, including small-
and large-scale farmers, processors, and consumers.
9.2 Developing sweetpotato farms as a niche
for conserving pollinators for multiple crops
In many cropping situations, sweetpotatoes are planted along
with other mass-produced crops including soybeans, corn,
and cotton. During flowering in the late summer, sweetpota-
toes can provide bees with valuable resources in a critical
period of summer when resources are scarce. Since sweet-
potatoes blossom during this time, the resources offered by
the flowers can be essential for bee populations in an envi-
ronment. However, sweetpotatoes are propagated vegetatively
and do not require pollination services for crop production,
creating little incentive for a sweetpotato producer to actively
maintain bee populations on their crop. Determining the
value sweetpotatoes bring to pollinators, which can improve
pollinator health and overall agricultural ecosystem produc-
tivity, can be a working point for developing shared interests
between growers and beekeepers. If sweetpotato cultivars are
used to help sustain pollinator populations, further research
will be needed to determine the possibility and extent of pol-
linators acting as vectors for sweetpotato pathogens (Real
et al., 2018). However, there can be advantages for incorpo-
rating pollinators, as honey bees can be used as a biomonitor
for hard-to-detect plant diseases (Cunningham et al., 2022;
Tremblay et al., 2019).
Floral resource nutrient profiles for sweetpotatoes need to
be established alongside other plants to strategically build a
landscape suitable for both growers and pollinators. The diver-
sity and health of different pollinators in sweetpotatoes are not
well understood and need to be compared with similar crop
landscapes without sweetpotatoes. These pollinators include
wild bees, which can be assessed for abundance and diver-
sity using passive and active sampling techniques. Using blue
vane traps and yellow pan traps, the diversity of bees and other
insects in the sweetpotato landscape can be sampled to get
more information on pollinators. From the grower’s perspec-
tive, honeybees are the most important managed pollinator
and can also be an informative biomonitor of sweetpotato dis-
eases. Nectar and pollen collected using front porch pollen

GEORGE ET AL.
traps can be analyzed for sweetpotato pathogens with targeted
approaches, such as PCR or ELISA assays.
9.3 Novel pest management tools and
disease detection methods
Higher costs associated with insecticidal applications for pest
management pose a financial burden on limited-resource
farmers and reduce profitability. In addition to the cost of
synthetic chemical insecticides, it adversely affects the envi-
ronment due to toxic buildup in the soil over time and is a
major cause of our food contamination. Socially disadvan-
taged small farmers are more vulnerable to losses due to
lack of IPM knowledge, limited resources, and challenges
in managing plant pests, as most IPM projects focus on
large farms (Collins, 2022). The polyphagous nature of wire-
worms and their long cryptic life cycles render chemical
control applications unsuccessful and pest estimation diffi-
cult under field conditions. Sweetpotato weevil pheromones
are already known and are used effectively for monitoring
weevils in multiple states. However, the semiochemicals for
click beetles (wireworms) on sweetpotato are unknown and
need further research. Identification of semiochemicals or
sex pheromones of click beetles can be effectively used for
sampling and monitoring wireworms in the field. Commer-
cially available biological insecticides and entomopathogenic
nematodes (EPNs) for the management of wireworms and
other insect larval stages on sweetpotato tubers need to be
evaluated. Laboratory, greenhouse, and field studies should
investigate the efficacy of EPN strains in controlling wire-
worms. Promising EPN strains should be used in further field
experiments for wireworm and root-knot nematode manage-
ment. Studies are required to investigate the soil microbial
community and the antagonistic effects of reducing parasitic
root-knot nematode incidence using EPNs.
Sweetpotatoes are prone to many viral and soilborne dis-
eases that can be vectored through the movement of infected
planting material. Producing disease-free planting materials
for growers, especially smallholder farms, is a big challenge.
Smallholder farmers often use heirloom seed materials that
have been passed on through generations, and very little is
known about the virus infection status of these planting mate-
rials. One solution to the problems caused by sweetpotato
“seeds” pathogens is to ensure that growers plant certified
clean propagation material. However, there is an estimated
72% shortfall of clean plant units needed to cover all sweet-
potato acreage (NCPN Network News, May 2018). Thus,
there is a considerable demand for establishing certified nurs-
ery farms to produce clean “seed” in the sweetpotato industry.
Besides, the high reinfection rate of viruses in production
requires the farmers to frequently purchase virus-indexed
propagation material, which greatly increases their financial
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21
input for propagation and production. Using high-throughput
sequencing, we can characterize the virus infection status
on those historical materials and identify the prevalence
of other soilborne pathogens in current “seeds” production
fields. By using different biotechnological approaches such as
gene transformation or genome editing techniques, disease-
resistant cultivars can be developed and are a promising
strategy to control SPVD viruses in sweetpotatoes.
9.4 Improving weed and water
management strategies in sweetpotato
cultivation
Weed management practices in smallholder and organic farms
are expensive and are the most critical obstacle to the adoption
of organic production or sustainable cultural practices (i.e.,
no tillage or minimum tillage). Adoption of cultivars toler-
ant to weed interference can be essential in integrated weed
management in conventional and organic production. Devel-
opment of erect and semi-erect vined sweetpotato germplasm
could allow increased tillage- inter row cultivation later into
the season, as the vines would not completely cover space
between the rows. Additional research is required to iden-
tify and develop sweetpotato genotypes that are fast growing
and have semi-erect to erect canopy architecture that may
provide other weed and insect management strategies for
sweetpotatoes.
Climate change is leading to increased extremes in droughts
and floods, in both frequency and intensity. Because of this
uncertainty in water supply, it is becoming more necessary
to understand the role water has in sweetpotato cultivation,
including WU, WUE, and runoff water quality and quantity.
Further research is required to measure and calculate WU
and WUE in the cultivar trials being conducted in the South-
eastern United States. Also, it is essential to measure runoff
quality and quantity at the onset of sweetpotato field experi-
ments. The findings from these studies can be used to inform
a hydrologic crop model to help simulate future manage-
ment possibilities. Different crop models have been reported
and evaluated for predicting future WU change from climate
variability. Some of these models have predicted reduced rain-
fall, thus modifying evaporation and runoff and indicating
an increased need for irrigation water. Similar crop mod-
els need to be developed for understanding WU, WUE, and
runoff in sweetpotato production for different cultivars and
geographical areas. Improving our knowledge on WU, WUE,
and runoff for sweetpotato production in North America
will allow for more informed management decisions leading
toward increased sustainability of production and water sup-
plies. Model testing and improvement with field experiments
would require a coordinated international effort and long-term
commitment to sweetpotato research.

22
10 CONCLUSIONS
In recent years, there is a growing demand and interest
in sweetpotato cultivation, and the number of projects and
scientists working on sweetpotato research has increased.
Smallholder farmers have realized that growing sweetpotatoes
is beneficial, and the increased demand may be due to the pro-
motion of the storage root’s health benefits. The sweetpotato
crop also has fewer insect pest problems than other storage
root crops, except for sweetpotato weevil and wireworms,
which cause significant damage in some of the sweetpotato
growing areas.
The current review demonstrates the need for research to
develop new sweetpotato genotypes and agricultural prac-
tices that maximize consumer sensory experiences and health
benefits. It may enhance the interest of small- and mid-
size organic farmers in adopting purple or other specialty
sweetpotato cultivars as a niche market crop. Advances in
sweetpotato chemistry and processing research will help
to add value and increase consumption. Novel sweetpotato
genetic materials with resistance to potyviruses will be devel-
oped by gene editing and can be used by smallholder farmers.
Germplasm evaluations will contribute to safeguarding and
utilizing the genetic diversity of the USDA sweetpotato
germplasm collection and its crop wild relatives in genetic
studies and breeding programs. EPNs that effectively control
wireworms and other insect larval stages will be identified
and used in smallholder and organic farms. A method of
using honeybees as an environmental biomonitor to detect
pathogens in sweetpotato cultivars will be developed. Iden-
tifying the antagonistic effects of EPNs and underground
semiochemicals involved will help to develop integrated man-
agement tools for smallholder farms. Increased knowledge
on WUE and runoff for sweetpotato production in North
America will allow for more informed management deci-
sions leading toward increased sustainability of production
and water supplies. Improving the irrigation water quality
and availability may increase yield and reduce pest incidence.
Using endophytic and mycorrhizal fungus holds promise in
increasing storage root yield and will be helpful for organic
and smallholder farms. The information provided in this com-
piled review also will help to enhance economic efficiency
and increase farm income, sustainability, and viability of
socially disadvantaged small farms.
AU T H O R C O N T R I B U T I O N S
Justin George: Conceptualization; investigation; method-
ology; supervision; visualization; writing—original draft;
writing—review and editing. Gadi Reddy: Conceptual-
ization; investigation; project administration; supervision;
writing—original draft; writing—review and editing.
Philip Wadl: Conceptualization; methodology; writing—
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GEORGE ET AL.
original draft; writing—review and editing. William
Rutter: Conceptualization; methodology; visualization;
writing—original draft; writing—review and editing.
Julianna Culbreath: Methodology; writing—original
draft. Pierre Lau: Conceptualization; methodology;
writing—original draft; writing—review and editing. Tahir
Rashid: Conceptualization; methodology; visualization;
writing—original draft; writing—review and editing.
Matthew Allan: Conceptualization; methodology; visu-
alization; writing—original draft; writing—review and
editing. Suzanne Johaningsmeier: Conceptualization;
methodology; writing—original draft; writing—review and
editing. Amanda Nelson: Conceptualization; methodology;
writing—original draft; writing—review and editing. Ming
Li Wang: Conceptualization; methodology; writing—
original draft; writing—review and editing. Augustine
Gubba: Methodology; writing—original draft. Kai-Shu
Ling: Conceptualization; methodology; writing—original
draft; writing—review and editing. Yan Meng: Con-
ceptualization; methodology; writing—original draft;
writing—review and editing. Daniel Collins: Conceptualiza-
tion; methodology; writing—original draft; writing—review
and editing. Sathish Ponniah: Conceptualization; method-
ology; writing—original draft; writing—review and editing.
Prasanna Gowda: Conceptualization; investigation;
methodology; project administration; resources; supervision;
writing—review and editing.
AC K N OW L E D G M E N T S
We would like to thank Shundalyn Moore for help with ref-
erences. The findings and conclusions in this publication are
those of the author(s) and should not be construed to represent
any official USDA or United States Government determina-
tion or policy. Mention of trade names or commercial products
in this publication is solely for the purpose of providing
specific information and does not imply recommendation or
endorsement by the United States Department of Agriculture.
C O N F L I C T O F I N T E R E ST STAT E M E N T
The authors declare no conflicts of interest.
ORCID
Justin George https://orcid.org/0000-0002-0807-5707
Ming Li Wang https://orcid.org/0000-0002-7139-1016
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